What Would Happen to the Trees and Lianas If Apes Disappeared?
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What would happen to the trees and lianas if apes disappeared? D AVID B EAUNE Abstract Apes, like many frugivorous animals, are crucial extended to the lesser apes, the gibbons (Hylobatidae; allies for the reproduction of several fruiting tree species. McConkey & Chivers, ). The bonobo was the last of Almost all apes, however, including bonobos Pan paniscus, the three African great ape species to be discovered. It is en- are threatened with extinction. How will this affect tree demic to the Democratic Republic of Congo, south of the conservation? How can plants that are adapted to seed dis- Congo River, and its habitat does not overlap with that of persal by apes reproduce without their dispersal vectors? At either chimpanzees or gorillas. Bonobos are mainly frugi- LuiKotale, in an evergreen tropical forest of the Democratic vorous (% of feeding sessions), spending c. hours Republic of Congo, the recruitment of plant species in the per day swallowing seeds (Plate ), which they transport absence of seed dispersal was investigated under the par- over a relatively long distance (% . m; mean . ental canopy, where a proportion of seeds fall without hori- km). An individual will disperse . t of seeds (. million zontal dissemination. Most bonobo-dispersed plant species seeds . mm in length, excluding those such as Ficus spp. (%of species) were unable to self-recruit under the can- and Musanga cecropioides) in its lifetime (Beaune et al., opy. As % of the tree species (% of trees) at LuiKotale b). More than plant species are dispersed by bonobos are dispersed by bonobos there is a risk of ecosystem decay through endozoochory (Beaune et al., b). Passed seeds and simplification (reduced biodiversity) if Pan paniscus germinate more rapidly, more successfully and with greater disappears from its natural range. The extinction of other post-dispersal survival than unpassed seeds (seeds embed- apes from their forests could have similar consequences. ded in faeces are involved in diplochory with dung beetles; The conservation of tree species, therefore, must encompass Beaune et al., a). In the forest of LuiKotale % of tree conservation of pollinators, seed dispersal vectors and other species and % of individual trees rely on bonobos for seed species that provide ecological services to the trees and other dispersal, and functional overlap between the bonobo and fruiting plants. other frugivorous animals appears to be low (Beaune et al., b,c). Keywords Africa, bonobo, Congo basin, ecosystem decay, The seed dispersal service provided by bonobos is of forest ecology, Pan paniscus, seed dispersal, zoochory particular interest as the species is categorized as Endangered on the IUCN Red List (Fruth et al., ), listed on CITES Appendix I (CITES, )andunder Class A in the African Convention on the Conservation of Nature and Natural Resources (OAU, ). The popu- Introduction lation is estimated to number ,–, (Fruth et al., ). The greatest threat to the bonobo is poaching for he conservation of a species is dependent upon the con- the commercial bushmeat trade but the species’ survival servation of those species that provide it with ecological T is also threatened by forest fragmentation and degra- services. The question of how the extinction of one species dation. Despite international protection and local taboos could affect others is particularly pertinent in the case of the bonobos are still killed in their natural habitats, including great apes (Hominidae: chimpanzee Pan troglodytes, bono- in protected areas such as Salonga National Park (Hart bo Pan paniscus, gorilla Gorilla gorilla and orang-utan et al., ). Pongo pygmaeus) because they play an important role as I assessed the ecological importance of bonobos for key dispersers of woody species (Wrangham et al., ; fruiting plants by investigating the ability of the plant com- Voysey et al., ; Gross-Camp & Kaplin, ; Beaune munity to recruit without horizontal seed dispersal (i.e. et al., a,b,c; Effiom et al., ). This can also be without bonobos and other seed dispersal vectors). The hy- pothesis is that without seed dispersal several plant species DAVID BEAUNE Department of Primatology, Max Planck Institute for for which apes are obligate seed dispersers would not re- Evolutionary Anthropology, Leipzig, Germany, and Laboratoire cruit under the parental tree because of density-dependent Biogéosciences, UMR CNRS 6282, Université de Bourgogne, Dijon, France E-mail [email protected] effects on the survival of seeds and seedlings. I also discuss – Received May . Revision requested July . how bonobos depend on trees in this plant animal Accepted October . First published online February . mutualism. Oryx, 2015, 49(3), 442–446 © 2015 Fauna & Flora International doi:10.1017/S0030605314000878 Downloaded from https://www.cambridge.org/core. IP address: 170.106.40.139, on 23 Sep 2021 at 21:26:18, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0030605314000878 Trees and apes 443 by bonobos (Beaune et al., b) were investigated during May –June , following the methodology of Chapman & Chapman (). All seedlings (, cm high), saplings (– cm high) and poles (. cm high, , cm diameter at breast height, DBH) were cen- sused in the parental crown area (i.e. the fruit-fall zone) of adult trees . cm DBH, with no conspecifics within m of the fruit-fall zone. The presence of seeds under the par- ental crown was confirmed by observation of falling seeds during animal feeding sessions and/or visual assessment of seeds present (fragments and fresh seeds). Mean numbers of seedlings, saplings and poles were calculated from a mini- mum of five adult plants per species. We considered an PLATE 1 Bonobo Pan paniscus eating velvet tamarind fruits adult plant to be able to self-replace when the mean pole (Dialium sp.) and swallowing seeds at LuiKotale, Democratic production per plant was $ . To confirm that a species Republic of Congo. Photograph: David Beaune/LKBP. was able to recruit outside its fruit-fall zone, density of re- cruits was estimated across the total census area. Results Nineteen plant species (three liana and tree species) were dispersed efficiently by bonobo (Table ). Three species con- sidered to be autochorous (i.e. dispersed by a plant’sown means) were included in the assessment for comparison (see Beaune et al., b,c for characterization of seed disper- sal). For each plant species some seeds were not dispersed horizontally and were found under the parental crown. During bonobo and monkey (Cercopithecus cephus ascanius, Cercopithecus mona wolfi, Lophocebus aterrimus)feedingses- sions broken branches and fruits with seeds (dropped or spat) from the fruiting plants were observed on the ground. The autochorous species recruited, on average, more than one pole under the parents, thus fulfilling the criterion for self- replacement. In contrast, the fleshy-fruited species dispersed by bonobos did not recruit enough poles under parent plants for self-replacement, except for Drypetes sp. (Table , Fig. ). FIG. 1 Location of the study site at LuiKotale, in Salonga Although seedlings, saplings and poles were found under National Park, Democratic Republic of Congo. other tree species (in a total of . ha censused), most adults of zoochorous (i.e. dispersed by animals) species were not able Study site to self-replace without seed dispersal beyond the fruit-fall zone, despite the presence of a seed bank. Fieldwork took place at LuiKotale, at the south-western fringe of Salonga National Park, Democratic Republic of Congo (Fig. ). The study site comprised . km of pri- Discussion mary evergreen lowland tropical rain forest (for more infor- The frugivorous bonobo relies on the fruits of plant spe- mation about the study site and previous research cies, which account for . % of their diet (Beaune et al., conducted there see Beaune et al., a,b,c, a,b,c,d,e). b). During certain months trees such as Dialium spp. can account for . % of the bonobo’s diet (Beaune et al., Methods d). How tree populations depend on the ecological ser- vices provided by bonobos is more difficult to assess. To assess seedling recruitment under the parent crown, ma- However, without seed dispersal, zoochoric plant species ture trees of species previously observed to fruit (thus ex- normally dispersed by bonobo recruit insufficient young cluding males of dioecious species) and efficiently dispersed under the fruit-fall zone for self-replacement, despite Oryx, 2015, 49(3), 442–446 © 2015 Fauna & Flora International doi:10.1017/S0030605314000878 Downloaded from https://www.cambridge.org/core. IP address: 170.106.40.139, on 23 Sep 2021 at 21:26:18, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0030605314000878 444 D. Beaune TABLE 1 Mean number of seedlings, saplings and poles found under the canopy of adult individuals of tree and liana species in LuiKotale, Democratic Republic of Congo (Fig. ). Mean recruitment Tree species1 (Family), limiting seed size2 (cm) No. of individuals Mean DBH (cm) Seedling Sapling Pole Autochory Hymenostegia mundungu (Caesalpiniaceae) 10 73.7 4.4 4.1 2.5 Scorodophloeus zenkeri (Caesalpiniaceae) 10 48.4 16.3 2 3.4 Strombosiopsis zenkeri (Olacaceae) 11 35.6 2 2.1 1.2 Zoochory with bonobo Pan paniscus Anonidium mannii (Annonaceae) ↔4 10 46.7 0.5 0.6 0.4 Blighia welwitschii (Sapindaceae) ↔2 5 62.8 0 0.2 0.4 Canarium schweinfurthii (Burseraceae) ↔1.2 5 109.4 0 0 0 Cissus dinklagei (Vitaceae) ↔1 5 0.8 0 0 Drypetes sp. (Euphorbiaceae) ↔1 10 30.9 0.7 1.9 2.6 Enantia olivacea (Annonaceae) ↔1 6 15.4 0 1.8 0.8 Ficus sp. (Moraceae) ø 0.1 7 0 0 0 Gambeya lacourtiana (Sapotaceae)↔ 1.9 10 92.2 1 0 0 Grewia oligoneura (Malvaceae) ↔2 6 38.4 0.2 0.3 0.3 Irvingia gabonensis (Irvingiaceae) ↔5 54 83.1 1.7 0.0 0 Irvingia grandifolia (Irvingiaceae) ↔5 10 110 0 0 0 Klainedoxa gabonensis (Irvingiaceae) ↔4 10 124.5 0 0 0 Landolphia forestiana (Apocynaceae) ↔1.2 5 0 0.2 0 Landolphia sp.